Berichte der Geologischen Bundesanstalt Vol 46-gesamt

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Berichte der Geologischen Bundesanstalt
Nr. 46

V International Symposium
Cephalopods -

Present and Past

Vienna
6 - 9th September 1999

Institute of Palaeontology, University of Vienna
Geological Survey of Austria
Museum of Natural History Vienna

ABSTRACTS VOLUME
Edited by
Kathleen Histon

Geologische Bundesanstalt
Vienna, July 1999

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Reference to this Volume:

HISTON, K. (Ed.)
V International Symposium Cephalopods - Present and Past, Vienna.
Abstracts Volume. Ber. Geol. Bundesanst. 46, 1-134, 111., Wien 1999
ISSN 1017-8880

Editor's address:
Kathleen Histon
Geological Survey of Austria
Rasumofskygasse 23
A-1031 Vienna
Austria

Impressum:
Alle Rechte für das In- und Ausland vorbehalten.
Copyright Geologische Bundesanstalt, Wien, Österreich.
Medieninhaber, Herausgeber und Verleger: Verlag der Geologischen Bundesanstalt,
A-1031 Wien, Postfach 127, Rasumofskygasse 23, Österreich.
Für die Redaktion verantwortlich: Kathleen Histon, Geologische Bundesanstalt
Layout: Kathleen Histon, Geologische Bundesanstalt
Druck: Offsetschnelldruck Riegelnik, A-1080 Wien
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Die "Berichte der Geologischen Bundesanstalt" sind im Buchhandel nur eingeschränkt erhältlich.

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Invited Speakers
Theo Engeser
Larisa Doguzhaeva

Berlin
Moscow

Scientific Board
Sigurd von Boletzky
Richard A. Davis
Larisa A. Doguzhaeva
Theo Engeser
Charles H. Holland
William J. Kennedy
Leopold Krystyn
Neil H. Landman
Walter L. Manger
Rudolf Schipp
Herbert Summesberger
Kazushige Tanabe
Gerd E.G.Westermann

France
USA
Russia
Germany
Ireland
UK

Austria
USA
USA
Germany
Austria
Japan
Canada

Organising Committee
Herbert Summesberger
Hans Peter Schönlaub
Leopold Krystyn
Kathleen Histon

Austria
Austria
Austria
Ireland

Sponsors:
Federal Ministry for Education and Cultural Affairs
"Austria Grant"
City Council of Vienna, Wissenschafts- und Forschungsförderung
"Vienna Grant"
Lord Mayor of Vienna, Dr. Michael Häupl
Town Council of Gumpoldskirchen
Geological Survey of Austria, Vienna
Museum of Natural History, Vienna
Institute of Palaeontology, Vienna
Acknowledgements

The staff of the Geological Survey of Austria, in particular Irene Zorn, Angelika Vrablik, Veronika
Zolnaritsch, Christian Widhalm and the members of the Palaeontology Department are thanked for their
generous help and support throughout the preparation for this conference. The editor is grateful to Albert
Daurer (Geological Survey) for his advice during the compilation of this volume. The assistance of the staff
of the Natural History Museum and Institute of Palaeontology was greatly appreciated.
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Contents
Part 1

ABSTRACTS LISTED IN THEMATIC ORDER

Part 2

AB STRACTS IN ALPHABETICAL ORDER OF FIRST AUTHOR
ADDRESS LIST OF AUTHORS

Parti
KEYNOTE PRESENTATIONS
DOGUZHAEVA, L.
Early shell ontogeny in bactritoids and allied taxa: comparative morphology, shell wall ultrastructure and
phylogenetic implication
32
ENGESER, T.
Phylogeny of the "Post-Triassic" Nautiloids

36

BIOLOGY OF RECENT AND FOSSIL CEPHALOPODS
ARKHIPKIN, A. & BIZIKOV, V.
Statolith shape as an indicator of the life style in recent and extinct decapod cephalopods

14

BEUERLE1N, K., GEBAUER, M., VERSEN, B. & SCHIPP, R.
Arterial hemolymph supply in the branchial hearts of the cuttlefish Sepia officinalis L. (Cephalopoda,
Dibranchiata)
20
BOLETZKY, S. VON
Yolk sac morphologies in cephalopod embryos

23

BOLETZKY, S. VON
Dwarf cephalopods: conditions of reproduction at small size

24

DAVIES, R.A. & MAPES, R.H.
Pits in internal molds of cephalopods

31

GLEADALL, I. G.
The terminal region of the male genital tract in cephalopod systematics: A revised terminology and an
illustration of intrageneric variation within one genus of Octopus
42

GONCALVES, I., SENDAO, J. & BORGES, T.C.
Octopus vulgaris (Cephalopoda: Octopodidae) gametogeneses: A histological approach to the verification of
the macroscopic maturity scales
44
HIELSCHER, B. & BOLETZKY, S. VON
Organization and reorganization of the main nerve cord in untreated and regenerating tentacles of the
cephalopod Sepia officinalis L
46
KENNEDY, W.J., COBBAN, W.A. & KLINGER, H.C.
Muscle attachment and mantle-related features in Upper Cretaceous Baculites from the United States Western
Interior
55
KEUPP, H.
Injuries - a key to understanding life modes of ammonoids

56

LANDMAN, N.H., KLOFAK, S.M., O'SHEA, S. & MIKKELSEN, P.M.
A giant squid in New York City

72

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LESCH, C , KITZROW, D., KÄMPFER, P., NEEF, A. & SCHIPP, R.
Microbiological and physiological studies on bacteria populations in the pericardial coelom of Nautilus

pompilius L. (Cephalopoda, Tetrabranchiata)
73
LEWY, Z.
The trophic control on the function of the Aulacoceratid and Belemnoid guard and phragmocone

74

MANGER, W. L., MEEKS, L.K. & RUSSELL, R.A.
Evaluation of septal crowding as an indication of sexual maturity in some Lower and Middle Carboniferous
ammonoids from the North American Midcontinent, United States
81
RICHTER, U. & FISCHER, R.
Soft tissue attachment structures on pyritized internal molds of ammonoids

95

ROUGET, I. & NEIGE, P.
Intraspecific variation of ammonoid embryonic growth stages and its bearing on post embryonic growth

97

RUTH, P. & SCHMIDTBERG, H.
Ciliated cells on the digital tentacles of Nautilus pompilius L. indicate their function as sense organs
(Cephalopoda, Tetrabranchiata)
98
SAUER, W. & MELO, Y.
Estimating actual fecundity of aloliginid squid

100

STEPHAN, D.A. & STANTON, R.J. Jr.
Impact of reproductive strategy on cephalopod evolution

106

SUMMESBERGER, H., JURKOVSEK, B. & KOLAR-JURKOVSEK, T.
Upper jaws of Placenticeratidae from the Karst Plateau (Upper Cretaceous, Slovenia)

107

TANABE, K. & LANDMAN, N.H.
Morphological diversity of the jaws of Cretaceous Ammonoidea

109

TANABE, K., MAPES, R.H., SASAKI, T. & LANDMAN, N.H.
Comparative microanatomy of the siphuncular cord in Permian ammonoids and Recent Nautilus

110

VAN DER TUUK, L.T. & JAGT, J.W.M
An enigmatic cephalopod jaw element from the latest Maastrichtian of the Netherlands

113

WESTERMANN, B. & SCHIPP, R.
Cytological investigations on the
Tetrabranchiata)

digestive

organs

of

Nautilus

pompilius

L.

(Cephalopoda,
119

CONSTRUCTION MORPHOLOGY
ARKADIEV, V.V. & VAVILOV, M.N.
Types of lobe line development of the Middle Triassic Ammonoidea of the Boreal Area

13

BÄNDEL, K.
Evaluation of constructional differences in Jurassic ammonite shells and Nautilus

17

CHECA, A.G. & KEUPP, H.
Regulation of coiling in planispiral ammonites, inferred from cases of infestation in vivo by epizoans

27

EBBIGHAUSEN, V., BECKER, R.T. & BOCKWINKEL, J.
Morphometric analysis of Paratornoceratinae (Goniatitida) from the early Famennian of southern Morocco... 35
KAPLAN, P.
Biomechanics as a test of functional plausibility: Testing the adaptive value of terminal-countdown
heteromorphy in Cretaceous ammonoids
54
KROGER, B.
Shell loss due to predation- effects on ammonoid buoyancy
5

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KULICKI, C , LANDMAN, N.H., HEANEY, M.J., MAPES, R.H. & TANABE, K.
Morphology of the early whorls of goniatites from the Carboniferous Buckhorn Asphalt (Oklahoma) with
aragonite preservation
68

KULICKI, C. & TANABE, K,
The ultrastructure of the dorsal shell wall of Mesozoic ammonoids

69

MAPES, R.H., TANABE, K. & LANDMAN, N.H.
Siphuncular membranes in Upper Paleozoic Prolecanitid Ammonoids from Nevada, USA

83

SPREY, A.
Early ontogeny of three Callovian ammonite genera (Binatisphinctes, Kosmoceras (Spinikosmoceras) and
Hecticoceras
105
WEITSCHAT, W.
Wrinkle layer and dorsal muscle scars in Amauroceras ferrugineum from the Domerian of NW Germany.... 118
WESTERMANN, G.E.G.
Recent hypotheses on mechanical and metabolic functions of septal fluting and sutural complexity in post120
Carboniferous ammonoids
YACOBUCCI, M.M.
Buckman's paradox: constraints on ammonoid ornament and shell shape

124

PHYLOGENY, SYSTEMATICS
AVRAM, E.
The taxonomic position and biostratigraphic value of the genus Pseudocrioceratites Egoian, 1969
(Lytoceratina) in Romania
15

BARDHAN, S., SARDAR, S. & JANA, S.
The Middle Jurassic ammonite Kheraiceras Späth from the Indian Subcontinent

18

BECKER, R.T.
Phylogeny and systematics of the Wocklumeriina (Ammonoidea, Clymeniida, Middle to Late Famennian).... 19
BOCKWINKEL, J., BECKER, R.T. & EBBIGHAUSEN, V.
Variability and taxonomy of Maeneceras (Goniatitida, Sporadoceratidae) from the early Famennian of
Southern Morocco

21
BOGDANOVA, T.N. & MIKHAILOVA, I.A.
Origin and evolution of the Family Deshayesitidae Stoyanow, 1949

22

DAGYS, A.
Phylogeny and taxonomy of the Boreal Anisian family Czekanowskitidae

29

DOGUZHAEVA, L.A., MAPES, R.H. & MUTVEI, H.
Rostrum and phragmocone structures in the lower Carboniferous coleoid Hematites and its taxonomic
assignment
33

DOGUZHAEVA, L.A., MUTVEI, H. & DONOVAN, D.T.
Structure of the pro-ostracum and muscular mantle in Belemnites

34

ENGESER, T.
The data retrieval system Nautiloidea (DRSN) + computer demonstration

37

GLOWNIAK, E.
Biology and biogeography of the middle Oxfordian ammonites of the subgenus Platysphinctes: a new evidence
from Poland
43

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HAAS, W.
The evolution of the Octopoda

45

HOEDEMAEKER, P.
Sexual dimorphism in the genus Pseudothurmannia

49

HOLLAND, C.H.
One of many problems: Taxonomy of the common Silurian Nautiloid cephalopod Orthoceras bullatum J. de C.
Sowerby
50
HOUSE, M.R. & BECKER, R.T.
The phylogeny of Pharciceratids and their relatives (Ammonoidea, Anarcestida:
Frasnian)

Late Givetian to Middle
51

HUDELOT, C. & BOUCHER-RODONI, R.
What molecular tools tell us about Octopods systematics

52

KORN, D.
Phylogeny of Early and Middle Devonian ammonoids

62

KOSTAK, M.
Coleoidea of the Bohemian Cretaceous Basin (Czech Republic, Europe)

64

KULLMANN, J.
Palaeozoic Ammonoidea in the database system Goniat + computer demonstration

71

KULLMANN, J.
Ammonoid evolution during the critical intervals before and after the Devonian-Carboniferous Boundary and
the mid-Carboniferous Boundary
70
MANGER, W.L., MEEKS, L.K. & RUSSELL, R.A.
Dimorphism in middle Carboniferous ammonoids from the southern mid-continent, United States

80

MUTVEI, H.
Nautiloid systematics based on siphuncular structure and position of muscle scars

89

NEIGE, P.
Diversity versus disparity: examples from present (Coleoids) and past (Ammonites) cephalopods

90

PLOCH, I.
Problem of sexual dimorphism in the Valanginian (Lower Cretaceous) ammonites Valanginites nucleus and
Saynoceras verrucosum
93
RODDA, P.U. & MURPHY, M.A.
Hypophylloceras and the classification of the Phylloceratidae

96

SERVENTI, P.
Some new Silurian nautiloid cephalopods from the Italian Carnic Alps

103

SERVENTI, P., GNOLI, M. & HISTON, K.
Revision of Silurian nautiloid cephalopods from the Carnic Alps from various museum collections

104

SZIVES, O.
Heteromorph ammonites from the Tata Limestone Formation (Lower Cretaceous, Aptian) Hungary

108

WARNKE, K., SÖLLER, R., BLOHM, D. & SAINT-PAUL, U.
Assessment of the phylogenetic relationship between Octopus vulgaris Cuvier, 1797 and O. mimus Gould,
1852 using mitochondrial 16SrRNA
117
ZEISS, A.
The Upper Jurassic ammonite fauna of Ernstbrunn (NE Austria) and its interesting position between Tethyian
and subboreal faunas
127

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TAPHONOMY, ECOLOGY, PALAEOECOLOGY
CHECA, A.G., HISTON, K. & SANDOVAL, J.
Cephalopod accumulations linked to condensation episodes in the Jurassic of the Subbetic (Southern Spain)
and in the Silurian of the Carnic Alps (Austria)
26
DALTON, R.B. & MAPES, R.H.
Scavenging or predation: Mississippian ammonoid accumulations in carbonate concretion halos around
Rayonnoceras (Actinocerida) body chambers from Arkansas
30
FERNANDEZ-LOPEZ, S., HENRIQUES, M.H. & DUARTE, L.V.
Taphonomy of Ammonite condensed associations. Jurassic examples from carbonate platforms of Iberia

38

FRAAYE, R.H.B.
Organisms in body chambers of fossil cephalopods

40

HISTON, K.
Telescoping in orthoconic nautiloids: an indication of high or low energy hydrodynamic regime ?

48

KLOFAK, S.M.
Size classes on a Devonian ammonoid from the Middle Devonian Cherry Valley Limestone of New York State,
USA
58
LUKENEDER, A.
Two ammonite mass-occurrences of the Alpine Lower Cretaceous (Northern Calcareous Alps, Upper
Austria)
75

LUKENEDER, A., HARZHAUSER, M., MANDIC, O. & ROETZEL, R.
Shell accumulation of the nautilidae Aturia (Aturia) aturi (Basterot, 1825) in the Retz-Formation (Lower
Austria, Upper Eggenburgian, Lower Miocene)
76
MACCHIONI, F. & PARISI, G.
Effects of compaction in ammonite moulds and its taphonomic implications: An example from the Rosso
Ammonitico of the Northern Appennines (Early-Middle Toarcian, Umbria-Marchean, Italy)
77
MANDA, S.
Wenlock and Lower Ludlow cephalopods in the Prague Basin: assemblages and palaeoecology

79

MAPES, R.H. & McCOMAS, G.A.
Septal implosion in coiled nautiloids from an Upper Carboniferous unit in Ohio, USA

82

MARCHAND, D., COURVILLE, P., SCOUFFLAIRE, Q., BONNOT, A. & ROSSI, J.
Ammonite faunas from marls with pyritic fossils (Lower Oxfordian): Original faunas at the interface distal
platform and basin
84
MARCINOWSKI, R.
Change within ammonite assemblages from Mangyshlak Mountains (Western Kazakhstan) during the MidCretaceous transgression
85
REBOULET, S.
Limiting factors on shell growth and segregation of ammonite populations: evidence from adult size variations
with time and space
94
SANDOVAL, J., O'DOGHERTY, L. & GUEX, J.
Sea-level, ammonite turnover and isotopic record in the early-middle Jurassic of the Betic Cordillera
(Spain)
99
SCHWEIGERT, G. & DIETL, G.
Preservation of cephalopods in the Upper Jurassic Nusplingen Lithographic Limestone (Late Kimmeridgian,
SW Germany)
101

TAVERNE, N., FRAAYE, R.H.B. & JÄGER, M.
Predation of early Jurassic ammonites

111
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TUREK, V.
Remarks to post-mortem behaviour of orthoconic shells

112

VÖRÖS, A.
Paleoenvironmental distribution of Middle Triassic ammonoids in the Balaton Highland (Hungary)

115

WIESE, F.
Middle Turonian to Lower Coniacian ammonite assemblages in Northern Germany, with reference to
Nostoceratids and Diplomoceratids
122

PALAEOBIOGEOGRAPHY AND STRATIGRAPHY
AGU1RRE URRETA, M.B.
Hemihoplitid ammonoids from the Austral Basin of Patagonia, Argentina

12

BALINI, M.
The classic Anisian (Middle Triassic) ammonoid localities of the Southern Alps and their significance for the
definition of the Anisian subdivisions
16
BONNOT, A., MARCHAND, D. & COURV1LLE, P.

Parallel biozonation in the Upper Callovian and the Lower Oxfordian based on the Sub-family Peltoceratinae
(Ammonitina, Aspidoceratidae)
25
CHRISTENSEN, W.K
Palaeobiogeography and migration in the late Cretaceous belcmnite Family Belemnitellidae

28

FERNANDEZ-LOPEZ, S. & MELENDEZ, G.
Trimarginia and Trimarginites (Ammonoidea) from the Iberian Basin

39

GALACZ, A. & MATIYA, B.
Passendorfer's middle Jurassic ammonites from the high Tatras

40 a

GAVRILOVA, V.A.
Ammonoidea of the Mangyshlak Lower Triassic

41

HILLEBRANT, A. VON
Paleobiogeography and relationship of South American Hettangian (Lower Jurassic) ammonites

47

JAGT, J.W.M.
Late Cretaceous ammonite faunas of the Maastrichtian type area

53

KLINGER, H.C. & KENNEDY, W.J.
Ammonoid palaeobiogeography - the Pseudoschloenbachia Paradox

57

KLUG, C.
Devonian ammonoid biometry and global events - preliminary results

59

KLUG, C. & KORN, D.
Ammonoid succession in Devonian Sections of Northwest Africa

60

KORN, D.
From Pliny to Walch - 1700 pioneering years of ammonoid research

61

KORN, D., PRICE, J. & HOUSE, M.
Ammonoid faunas from the Devonian and Early Carboniferous of the Carnic Alps

63

KRYSTYN, L.
A major Phylloceratid - Lytoceratid faunal turnover in the Lower Jurassic

66

KRYSTYN, L. & BALINI, M.
Triassic ammonoids and the Himalayas

67

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MACHALSKI, M.
The last Maastrichtian ammonites in Poland.

78

MATYJA, B. & WIERZBOWSKI, A.
Biological response of ammonites to changing environmental conditions: An example of Boreal Amoeboceras
invasions into Submcditerranean Province during Late Oxfordian
86
MORIYA, K. & HIRANO, H.
Ammonoid assemblages in the Santonian of Hokkaido, Japan, with special reference to the Desmoceratid
ammonoids
87
MOUTY, M. & GAUTHIER, H.
Mid-Cretaceous ammonites from the coastal chain of Syria

88

PÄLFY, J.
Ammonoid biostratigraphy of the Triassic-Jurassic Boundary near Csövär, Hungary: A progress report

92

PÄLFY, J.
Early Jurassic Ammonoids from the Persani Mts. (East Carpathians, Romania)

91

SEIBERTZ, E. & SPAETH, C.
Range and distribution of belemnitcs in the Jurassic and Cretaceous of Mexico

102

VASICEK, Z
Palacobiogeography of the early Cretaceous (Prc-Aptian) ccphalopod bearing formations of the Western
Carpathians (Czech and Slovak Republics) and the Northern Calcareous Alps (Austria)
114
VÖRÖS, A.
Triassic ammonoids and bioslratigraphy of the Balaton Highland: New results from the Anisian, Ladinian and
Carnian
116
WESTERMANN, G.E.G
Cephalopods, Vienna: Birth of marine Palcobiogcography

121

WILMSEN, M.

Upper Cretaceous nautiloids from northern Cantabria, Spain

123

YAZYKOVA, E.A.
Campanian-Maastrichtian ammonites from Far Eastern Russia, stratigraphy and palaeobiogeography

125

YUN CHEOL - SOO
Biogeographical characteristics of the Ordovician cephalopods from Korea

126

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Part 2

ABSTRACTS

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HEMIHOPLITID AMMONOIDS FROM THE AUSTRAL BASIN OF PATAGONIA, ARGENTINA
Aguirre-Urreta, Maria Beatriz

Departamento de Ciencias Geologicas, Universidad de Buenos Aires, CONICET.

The recent finding of a beautifully preserved ammonite fauna in the Austral basin of Southern Patagonia,
Argentina sheds light on the origin, evolutionary trends, and palaeobiogeographic distribution of the genus
Hemihoplites Späth, 1922.
Two species were already known from Patagonia: Hemihoplites ploskiewiczi and H. varicostatus. The former is
a small species with subrectangular to compressed whorl and fine, flexuous ribbing and has been included in the
Favrella americana assemblage zone (late early to early late Hauterivian). The second one is a medium sized
species with subrectangular to subquadrate whorl, flexuous ribbing bifurcate in inner whorls and simple on outer
whorls. It has been found in beds with Protaconeceraspatagoniense {Favrella wilckensi assemblage zone, late
Hauterivian) and Hatchericeras spp. (H. patagonense assemblage zone, early-middle Barremian) (Riccardi and
Aguirre-Urreta, 1989). Both species show sexual dimorphism. A third species, reported here for the first time,
corresponds to Hemihoplites feraudianus (d'Orbigny), type species of the genus. Several specimens recovered
from a single bed shows a total coincidence with the modern descriptions of this species regarding size, whorl
section, and ornamentation (Delanoy, 1990,1997). This species is also represented by a dimorphic pair. No other
ammonites have been recorded with this species, but it can be placed in the Late Barremian (Feraudianus zone
of Europe).
Hemihoplites is recorded in the recent edition of the "Treatise" as Upper Hauterivian-Barremian and its presence
in older rocks of Southern Patagonia as recorded by Riccardi and Aguirre-Urreta (1989) has been overlooked
(Wright, 1996) or dismissed as unlikely (Delanoy, 1990).
The early appearance of Hemihoplites in Southern Patagonia led some authors to question the age of the Favrella
americana, F. wilckensi and Hatchericeras patagonense faunas (Delanoy, 1990). However, the recent discovery
of Hemihoplites feraudianus permits to postulate a long history of this genus in the Austral basin, with an
evolutionary trend towards increasing body size and coarser ribbing.
The sudden appearance of Hemihoplites feraudianus in the European late Barremian also prompts to the proposal
of an immigration to the Tethyan region and casts doubts on the proposed origin of Hemihoplites from the late
Hauterivian genus Pseudothurmannia (Wright, 1996) or from Emericiceras of the barremense group (Delanoy,
1990).

DELANOY, G.,

1990. Donnees nouvelles sur l'espece-index Hemihoplites feraudianus

(d'Orb.,

1841)(Ammonoidea, Ancyloceratina). - CR. Academie Sciences Paris 310, Serie II, 661-666.
DELANOY, G., 1997. Biostratigraphie des faunes d'Ammonites a la limite Barremien-Aptien dans la region
d'Angle-Barreme-Castellane. - Annales du Museum d'Historie Naturelle de Nice, 12, 1-270.
RICCARDI, A.C. and M.B. AGUIRRE-URRETA, 1989, Hemihoplitid ammonoids from the Lower Cretaceous
of Southern Patagonia. - Palaeontology, 32,447-462.
WRIGHT, C.W., 1996. Cretaceous Ammonoidea. In: R.L. Kaesler: Treatise on Invertebrate Paleontology, Part
L, Mollusca 4 Revised , 362 pp. Geological Society of America and University of Kansas Press.
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TYPES OF LOBE LINE DEVELOPMENT OF THE MIDDLE TRIASSIC AMMONOIDEA OF THE
BOREAL AREA
Arkadiev V.V.*. VavilovM.N.**
*Russia, 199026 Saint Petersburg, Mining Institute Russia, 191104 Saint Petersburg,
**All-Russia Petroleum Research and Geological Prospecting Institute

Four-lobed (VL:ID) or, more rare, five-lobed (VLU:ID) prime suture has been established by the authors to
be characteristic for the majority of the Middle Triassic ammonoidea of Boreal region. Representatives of
families Longobarditidae, Nathorstitidae, Tsvetkovitidae, Parapopanoceratidae, Ptychitidae, Beyrichitidae,
Ussuritidae have prime sutures of the first type, Proarcestidae and Cladiscitidae - of the second type. Main
types of lobe line development of the Middle Triassic Ammonoidea are:
1. Bilateral displacement of umbilical lobes, with the even lobes being displaced to the external side, the odd
lobes - to the internal side of the whorl. Typical formula is (V,Vi)LU1U2U4..U3I(DiD1). Families

Longobarditidae, Nathorstitidae and Tsvetkovitidae.
2. Bilateral displacement of umbilical lobes, with the odd lobes being displaced to the external side, the even
lobes - to the internal side of the whorl. Typical formula is (V1V,)LU1U3..U2I(DiD1). Family
Parapopanoceratidae.
3. Formation of umbilical lobes on the external side of the whorl and additional internal lateral lobes - on the
internal side of the whorl, according to the formula: (ViV1)LU1U2U3..II2I1(D]D1). Family Ptychitidae.
4. Fission of the internal lateral lobe (I) on the early and middle ontogenesis stages after formation of the first
umbilical lobe (U1) on the external side of the whorl. Typical formula is CV^VOLU'l^LXDiD]). Family
Ussuritidae.
5. The lobe line is developed according to the fourth type, but new lobes Ii, I2, I3 etc. are forming from the
saddle dividing the wings of the lobe (I) after its fission. Typical formula is (ViVi)LU1IvI2I3-.-I4I1Id(DiDi).
Family Beyrichitidae.
6. A peculiar type of the lobe line development has been discovered for the Middle and Late Triassic
Arcestidae having five-lobed prime suture. The development of the genus Nevadisculites studied by the
authors from the Middle Anisian of Nevada occurs primarily by fission of the lobe (I) followed by formation
of umbilical and additional internal lateral lobes, with first of them being displaced to the external side and
the second - to the internal side of the whorl according to formula: VL:U:ID - VLU:ID -VLU^iIiD (V]V1)LUU1U2:(I1.2Ii.2)Iil'(DiDl). Analogous type of development is characteristic for the Carnian
Proarcestes. The only distinction is that an adventive lobe occurs at the early stage (0.8 - 1 whorl) on the top
of the saddle V/L and disappears at the stage of 1.3 - 1.5 whorls. Both genera Nevadisculites and Proarcestes
are included by the authors into the Family Proarcestidae. It is suggested that some Triassic Ammonoidea,
due to the presence of five-lobed prime suture, should be excluded from the order Ceratitida having fourlobed prime suture. Triassic Ammonoidea having five-lobed prime suture join into the order Arcestida
Vavilov, 1992.

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STATOLITH SHAPE AS AN INDICATOR OF THE LIFE STYLE IN RECENT AND EXTINCT
DECAPOD CEPHALOPODS

Arkhipkin Alexander*. Bizikov Vyacheslav **
"Fisheries Dept., Stanley, Falkland Islands (fish.ßßc)**VNIRO, 17a V.-Krasnoselskaya street,
Moscow, 107140 Russia ()

Squids and sepioids are decapod cephalopods having the unique jet propulsive locomotion which is
characterized by a rhythmic change of linear acceleration. We have found that under acceleration, the
statolith (gravity stone) may deviate around the three mutually perpendicular axes running through its
rotation center and therefore may induce endolymph flows within the cavity of the equilibrium organs
(statocysts). This finding re-considers the theory of the statocyst function in decapods. As the statoliths play
an important role in detection of various types of accelerations, their general morphology is different in
pelagic and demersal decapods (Fig.l). Pelagic squids (both nektonic and planktonic species) live in an
environment without physical obstacles, and therefore have mainly 'rocket-like' movement. Demersal
sepioids and near-bottom squids either have to land on or
to take off from the bottom, and negotiate bottom
obstacles. Peculiarities in the statolith shape and size of
demersal decapods (their relative total statolith length is
considerably greater than that of pelagic species) make
their statoliths more movable around the transversal axis
than those of pelagic squids. Such a mobility provides a
greater sensitivity to accelerations of the animal during
both gliding and pitching. The oar-like rostrum of the
demersal decapod statolith provides a greater sensitivity
during rolling at low accelerations. Distinct separation of
the statolith body into dorsal and lateral domes provides
separation of endolymph flows in the corresponding
channels during pitching and yaw, respectively. Thus,
peculiarities of the statolith shape in demersal decapods
Fig. 1. Statoliths of the near-bottom squid
Berryteuthis magister (A) and pelagic squid
Moroteuthis robusta (B). Lateral (left) and

anterior (right) views. Dorsal (Dd) and lateral
(Ld) domes, dorsal spur (Ds), statolith body
(Sb), spur (S), wing (W) and rostrum (R).

give

them

greater

sensitivity

to

low

angular

accelerations in all possible planes compared to those of
pelagic species. Peculiarities of the statolith shape in
pelagic species (especially

long and wide wing

achieving the rostrum tip and short sharpened rostrum)
make these statoliths hardly movable around both longitudinal and transversal axes that considerably
diminish the sensitivity of these animals to angular accelerations in pitching and rolling planes. In pelagic
squids, the statolith monitors mainly the strength and frequency of jet propulsions during linear movement.
Thus, our findings make it possible to re-construct the life style and movement patterns of recent and (most
important) extinct decapods using statolith shape features, and to re-consider evolutionary connections

between different decapod groups.
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THE TAXONOMIC POSITION AND BIOSTRATIGRAPHIC

VALUE

OF THE

GENUS

PSEUDOCRIOCERATITES EGOIAN, 1969 (LYTOCERATINA) IN ROMANIA
Avram, Emil
Geological Institute of Romania, 1 Caransebes str., R-78344 Bucharest, Romania.
Email:

In describing his genus Pseudocrioceratites, Egoian (1969, p. 172) stated that it includes only two species:
P. pseudoelegans EGOIAN and P. rotundus EGOIAN, both of them Clansayesian in age. Nevertheless, this
group of cephalopods, characterised by a crioceratic appearance, but with typical lytoceratic suture, is also
represented in the Barremian (by two new Romanian species and, possibly, by the incompletely defined
Pseudocrioceras stentor ANDERSON), in the lower Aptian (by another Romanian species), and in the
Albian time span (by Crioceras cf. Munieri DOUVILLE, 1916, non SARASIN & SCHONDELMAYER,
1901).

In this acception, the genus displays a very near to Crioceratites emerici LEVEILLE costume (except for the
septate tubercles), and the suture with a very large, bifid lateral lobe and an also large, tending to be
cruciform, internal lobe. On this basis, the genus is here interpreted as being an uncoiled representative of

typical Lytoceratina, similarly to the related genus Acantholytoceras SPÄTH, 1923.

References.

EGOIAN, V.L., 1969: Ammonity iz klansejskikh sloev Zapadnogo Kavkaza. - Trudy Vses. Neftegaz.
Nauchno - issled. In - ta, Moskva, 19, 126-188 (In Russian)
DOUVILLE, M.H., 1916: Les terrains secondaires dans le Massif du Moghara a Test de l'Isthme de Suez,
Paleontologie. - Mem. Acad. Sei. Paris, ser. 2, 54, 1-184.

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THE CLASSIC ANISIAN (MIDDLE TRIASSIC) AMMONOID LOCALITIES OF THE SOUTHERN
ALPS, AND THEIR SIGNIFICANCE FOR THE DEFINITION OF THE ANISIAN SUBDIVISIONS
Balini Marco
Dipartimento di Scienze della Terra, Via Mangiagalli 34, 20133 Milano, Italy.
E-mail:

The development of the standard scale for the Anisian Stage (Middle Triassic) has been a matter of debate
since the last century. The Anisian Stage was defined in the Western Tethys, and its meaning and subdivisions
were strongly influenced by ammonoid rich localities of Southern and Northern Alps, as well as Hungary.
Many problems came out because of: (1) lack of bed-by-bed data, (2) restricted stratigraphic extent and
scarce overlap of some type-localities, or (3) historical localities with condensed faunas, (4) direct
correlations influenced also by taxonomical problems. For these reasons the number and the composition of
the biozones of the Middle to Upper Anisian (Pelsonian and Illyrian Substages) is not yet well defined.

In the last 10 years seven classical and new localities in the Southern Alps have been bed-by-bed investigated
in order to define the details of the Pelsonian-Illyrian faunal succession. The studied sections include Dont

(Dolomites), Cimego (Giudicarie) that are the two type localities of the Pelsonian Substage and of the
Binodosus Zone, and Stabol Fresco section, that is the most representative for the Illyrian Substage and the
Trinodosus Zone in the original definitions. Adanä and Peschiera (Giudicarie), Menna and Lenna (Brembana
Valley) were also taken into consideration.

All the sections were bed-by-bed directly correlated by guide ammonoids. In particular the most important
result is the new direct correlation of Dont and Stabol Fresco sections.

The first ammonoid level of Stabol Fresco section, providing a rich Bulogites fauna, is correlated with level R
of Dont section, that is some 2 m below the classic binodosus fauna. New fossiliferous levels have been found
at Dont 2.5 m above the layers providing the binodosus fauna. The new fossil bearing beds record the FO of
the bivalve Daonella followed by the occurrence of Megaceratites, Lanceoptychites and Paraceratites s.s.
These taxa, typical of the trinodosus fauna, occur in the same sequence in the middle-upper part of Stabol
Fresco section, some 50 m above the Bulogites-rich level. At Stabol Fresco, in between the last Bulogites
level and the typical trinodosus fauna there is a rich fauna dominated by ceratitids of the group of C.
cimeganus Mojsisovics, 1882. This fauna is not recorded at Dont where at the top of the binodosus-b&ax'mg
level a lithologic change underlined by glauconite suggests a hiatus. Moreover faunal analysis of the
binodosus-bearing leads to suggest a condensation.

The cross correlation of the studied sections are summarized in a composite range chart that notably differs
from recent literature. As a consequence a new biostratigraphic scale for the Pelsonian-Illyrian is proposed.

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EVALUATION OF CONSTRUCTIONAL DIFFERENCES IN JURASSIC AMMONITE SHELLS
AND NAUTILUS
Bändel, Klaus

Geologisch-Paläontologisches Institut und Museum, Hamburg.

Modern Nautilus has served as a model in understanding the construction and function of the ammonite
phragmocone. In the early Upper Jurassic of Madagascar a nautiloid that is essentially like modern Nautilus
occurs together with the representatives of the three orders of the ammonites, Lytoceratina, Phylloceratina,
and Ammonitina. All these are preserved without changes in their mineral shell structure, and in part, with
their siphuncular tube well preserved. The different features of shell that resulted from septum formation,
chamber sheet attachment, siphonal-tube construction, muscle attachment and internal living chamber
coatings (wrinkle layer, callus layer) are well preserved, while some organic layers have disappeared during
early diagenesis, that resulted in no shell crushing due to compaction. The features interpreted in the light of
new data that we have collected in our most recent studies on chambered cephalopods in Hamburg, Berlin,
and Hanover give new insights into ammonite paleobiology.

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THE MIDDLE JURASSIC AMMONITE KHERAICERAS SPÄTH FROM THE INDIAN
SUBCONTINENT
Bardhan. Subhendu. Sardar, Subrata, Jana, Sudipta
Department of Geological Sciences, Jadavpur University, Calcutta-700032,India
E-mail: subhendu ©jugeo.clibO.ernet.in

Kheraiceras Späth has a near circumglobal distribution except for subboreal and is marked by a 'bio-event'
(radiatio) especially during the Late Bathonian and Early Callovian time. Such 'bio-events' may blur the
distinction of faunal provincialism and cut across geographic boundaries and thus help in establishing
regional standard chronostratigraphy and interprovincial correlation. The present study reports six
Kheraiceras species from the Indian subcontinent (Kutch, India and Baluchistan, Pakistan) including three
new species. They are Kheraiceras cosmopolitum (Parona and Bonarelli), Kbullatum (D'Orbigny,), K.cf.

hannoveranum (Roemer), K.spathi n.sp.A and K.noetlingi n.sp.
Biostratigraphic potentialities of Kheraiceras are also explored. Dimorphism is well understood in the genus;
but specific dimorphic pairs are poorly known. At least in three instances, matching of pairs has been firmly
established and two new microconchs and one macroconch still lack their partners. Microconchs are of great
help in species discrimination, biozonation and understanding evolution.

18


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PHYLOGENY

AND

SYSTEMATICS

OF

THE

WOCKLUMERIINA

(AMMONOIDEA,

CLYMENIIDA, MIDDLE TO LATE FAMENNIAN)
Becker, Ralph Thomas
Museum für Naturkunde, D-10115 Berlin,

In one of his classic monographs, Schindewolf (1937) described the ammonoid biostratigraphy of the latest

Famennian (Wocklum-Stufe, Upper Devonian VI) which is based to a large extent on the rapid evolution of
clymeniids with strange triangular coiling, also known, for example, in some Carboniferous goniatites. Trigonal
whorls were thought to have evolved in three parallel lineages, two of which terminated abruptly during the
sudden mass extinction associated with the global Hangenberg Event.
Since Schindewolf's work, new rich faunas with triangular clymenids have been described from SW England,
Algeria, South China, Russia, Kazakhstan, Thuringia, and Poland. These records, a revision of German
collections (> 1.500 specimens), and new collections from Morocco, the Rhenish Massive, and Oklahoma
allow a re-evaluation of systematics and phylogeny. Neo- and lectotypes had to be selected for all German
taxa and there are new species of Synwocklumeria and Kielcensia. Intermediate Russian and Polish taxa show
that Wocklumeriidae with ventral lobe were derived from open umbilicate Parawocklumeriidae (Korn 1992).
Both groups form a natural systematic unit (Wocklumeriaceae) characterized by three apomorphies: 1.
triangular coiling of at least early stages, 2. weak to strong constriction of straight whorl parts, and 3. the
dorsal lobe is divided or reduced. Heterochronic processes played a major role in phylogeny. Parawo.
paradoxa,

Epiwo. applanata,

Wo. sphaeroides,

and Kielcensia bohdanoviczi represent

iterative

hypermorphotic end-members, partly with re-activated ancestral conch features such as evolute and rotund
coiling. Intraspecific dimorphism in the latter two species is documented by micromorphic specimens with
shortened triangular ontogenetic stages. Heterochronic speciation (miniaturisation) may have been in progress
just before the sudden extinction caused by the Hangenberg anoxic event. Large intraspecific variability of
shell parameters in some species, slowly expanding longidomic whorls, deep protective constrictions, and the
reduction of the ocular sinus indicate a suprabenthonic (demersal) lifestyle of Wocklumeriaceae. Triangular
coiling can be interpreted as an attempt to shift the centre of mass and aperture downwards in order to

optimize seafloor feading during long phases of growths.
Shells features and sutural patterns support Schindewolfs idea of an origin of Wocklumeriaceae in
Hexaclymeniidae although both groups are separated by a record gap. Glatziellidae, Pachyclymeniidae and
Biloclymeniidae are regarded as further hexaclymeniid side-branches. All clymenids with primarily wide and
later subdivided ventral lobe are assigned to the suborder Wocklumeriina. Gonioclymenids with narrow ventral
lobes were derived from Platyclymeniidae. This confirms the dissolution of the traditional Gonioclymeniina
(Korn 1992) and requires a re-definition of the Clymeniina.

Korn, D. (1992): Relationship between shell form, septal construction and suture line in clymenid cephalopods
(Ammonoidea; Upper Devonian). - N. Jb. Geol. Paläont., Abh. 185, 115-130.
Schindewolf, O.H. (1937): Zur Stratigraphie und Paläontologie der Wocklumer Schichten (Oberdevon). - Abh.
preuß. geol. L.-Anst., n.F. 178, 141 pp.
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ARTERIAL HEMOLYMPH SUPPLY IN THE BRANCHIAL HEARTS OF THE CUTTLEFISH
SEPIA OFFICINALIS L. (CEPHALOPODA, DIBRANCHIATA)
Beuerlein, Knut, Gebauer, Martin, Versen, Bernhard, Schipp, Rudolf
Institut für Allgemeine und Spezielle Zoologie, Justus-Liebig-Universität, 35390 Giessen, Germany;
Laboratoire d'Oceanographie Biologique, Universitö de Bordeaux, 33120 Arcachon, France
The circulatory system of dibranchiate cephalopods includes the systemic heart, the paired branchial hearts
and autonomously contractile vessels (Fiedler & Schipp, 1987; Wells & Smith, 1987). In the respiratory
part of the venous system mainly the branchial hearts are responsible for pumping the deoxygenated hemolymph into the gills. There, the oxygenation of the respiratory pigment hemocyanin takes place. In octopods
the average oxygen content in the arterial hemolymph is 3.3 vol %, whereas in the dichotomous branching
of the vena cava, which supplies the branchial hearts with hemolymph, the oxygen content is only 0.4 vol %
(Johansen & Lenfant, 1966) and may approach zero during hypoxic excursions (Houlihan et al., 1982).
Concerning the moderate oxygen supply of the branchial heart Driedzic (1985) proposed that it primarily
operates anaerobically. However, the demonstration of arterial vessels running to the muscular rind of the

branchial heart of Octopus (Wells & Smith, 1987), and the comparable enzyme profile of systemic and
branchial heart (Driedzic et al., 1990), indicate an independent oxygen supply of this organ.
For the further elucidation of the existence of an arterial vascularization in the branchial hearts of Sepia
officinalis L., tracer experiments were carried out on semi-adult animals of both sexes caught in the Bassin
d'Arcachon (France). After in-situ injection of 1 ml Indian Ink (0.1% in seawater) into one of the auricles,
the capacious organ complex composed of systemic heart, auricles, gills, renal appendages, branchial hearts
and branchial heart appendages was removed. Tissues were investigated by histological methods.
Marked vessels coming from the connective tissue at the base of the branchial hearts were mainly localized
at the dorsal surface of the organs by macroscopical observations. They form a ramifying mesh directly
beneath the epithelium; some of these vessels run through the muscular wall up to the branchial heart
appendages. The light microscopical studies revealed that the vessels of the inner folded epithelium in the
branchial heart appendages are also filled up with tracer particles. In the extended lumen of the branchial
hearts no tracer was found. After control injection of the Indian Ink solution into the vena cephalica, tracer
particles were detected in the lumen of the organs but not in the ramifying vessels mentioned above.
The presented morphological study verify that the branchial hearts and even their appendages are supplied
with oxygenated hemolymph from the systemic heart via a well-developed arterial vascularization - similar
to the coronary system as described for the heart of most vertebrates.

Driedzic, W.R., 1985: Journal of Experimental Biology, 117,471-474
Driedzic, W.R., Sidell, B.D., Stewart, J.M., Johnston, I.A., 1990: Physiological Zoology, 63, 615-629
Fiedler, A., Schipp, R., 1987: Experientia, 43, 544-553
Houlihan, D.F., Innes, A.J., Wells, M.J., Wells, J., 1982: Journal of Comparative Physiology, 148, 35-40
Johansen, K., Lenfant, C, 1966: American Journal of Physiology, 210,910-918
Wells, M.J., Smith, P.J.S., 1987: Experientia, 43,487-499
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VARIABILITY AND TAXONOMY OF MAENECERAS (GONIATITIDA, SPORADOCERATIDAE)

FROM THE EARLY FAMENNIAN OF SOUTHERN MOROCCO
Bockwinkel. Jürgen1, Becker, Ralph Thomas2, Ebbighausen, Volker3
('Dechant-Fein-Str. 22, D-51375 Leverkusen, 2Museum für Naturkunde, Invalidenstr. 43, D-10115 Berlin,
, 3Engstenberger Höhe 12, D-51519 Odenthal; all Germany)

The Sporadoceratidae are one of the most succesful and longest-ranging Famennian ammonoid family with
global distribution in (sub-)tropical areas. At the base of the classical Upper Nehden-Stufe (do Ilß = Upper
Devonian II-G), the oldest genus of the family, Maeneceras, descended from Cheiloceras (Puncticeras)
lagowiense by the insertation of shallow lobes in the ventral (A2-lobe) and dorsal (U-lobe) saddles. More than a
dozen early Famennian (Nehden-Stufe) species have been named which are said to be distinguished by shell
parameters, constrictions (internal shell thickenings), and by the shape and depths of sutural elements. Similar
species (Maeneceras ungeri Group) occur higher in the Famennian (UD III-V) but are all characterized by
spiral ornament as in many Carboniferous goniatites and they will be placed in a new genus.

In order to clarify the taxonomic status and variability of early maeneceratids, a large population (nearly 300
specimens) from a single bed low in UD II-G at Hassi Nebech (southern Tafilalt, Anti-Atlas) has been used for
morphometric investigation. Apart from usual shell parameters (whorl height and width, apertural height), the
number and extent of mould constrictions per whorl and the relative proportions between the two adventitious
lobes have been measured. There is moderate variation in whorl expansion rates but larger variation can be
seen in whorl width. A number of specimens reach values as in the lectotype of M. rotundum (Wedekind) but
there is only a slight separation of thicker and more compressed specimens.
The analyses of mould constrictions confirms the former idea that these follow growth stages with distinctive
patterns. From 10 to 50 mm diameter, the number of constrictions increases constantly from between 2 and 3 to
between 5 and 8 per whorl. In early stages, constrictions are complete but they gradually terminate on the lower
or outer flanks in subsequent whorls. From ca. 60 mm diameter, constrictions are reduced drastically which
suggests that protection against shell-breaking predators became irrelevant at late maturity. Clearly, taxa based
on the number and course of constriction only, cannot be regarded as valid.

Previous studies showed that the second A-lobe became deeper during phylogeny, culminating in species such
as M. milleri and M. acutolaterale where the rounded A2 is as deep or deeper than A]. The investigated

assemblage shows no relationship between size and depth of A2 which suggests that forms with various degrees
of septal folding were present contemporaneously. A histogram displays a small group with extremely shallow
A2 sensu M. nuntio, a majority group with A2/Ai ratio between 16 and 28 % as in M subvaricatum, and a
slightly separated second peak between 32 and 40 % (trend towards M. latilobatum). The population studied
gives no clear distinction of biological species; continuing morphological separation in succesive faunas yet has
to be shown. However, the subsequent introduction of additional morphological features such as spiral
ornament only in the main group with Ai < 30 % A2 leaves the possibility that the single bed studied documents
an episode of maeneceratid speciation in progress.
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THE ORIGIN AND EVOLUTION OF THE FAMILY DESHAYESITIDAE STOYANOW, 1949
Bogdanova Tamara N.1, Mikhailova Irina A.2
'All-Russian Geological Institut (VSEGEI), St.-Petersburg, Srednyi pr., 74, 199106, Russia, e-mail:
2Moscow University, Moscow, Leninskie Gory, 117234, Russia

The family Deshayesitidae represents the important stage of the Early Cretaceous ammonite evolution. The
subdivision of the Lower Aptian deposits practically in all the continents of the Earth is based on
interrelations within some genera and species associations of this family. Morphology of the shell and
evolution of suture of the European, Caucasian and Turkmenian deshayesitids have been revised. As a result,
we propose the new concept of some generic taxa of the Deshayesitidae. Deshayesites Kasansky, 1914 rather compressed, with flat or slightly convex flanks, subtriangular sections with a broad arched venter,
strong ribs, sigmoidal primaries and branching or intercalated secondaries (1 or 2). No distinct tubercules.
Suture formula - VUII^l'D: loben I, I2 - on the flanks. Type-species: Ammonites deshayesi (Leymerie in
d'Orbigny, 1842), Lower Aptian, France (Paris Basin). It includs about 40 species. Paradeshayesites
Kemper, 1967 - different size of shell, very involute, whorls high, triangulate with flat flanks and arched
venter; ribs dense, fine with bullates in the base and bundles, primaries bifurcate and trifurcate, secondaries 4
to 7-9. Suture formula - VUII 2 1 3 :l' D, loben I, I2 - on the flanks. Type-species: Hoplites laeviusculus (von
Koenen, 1902), Lower Aptian, tenuicostatus Zone, North-West Germany. It includes about 20 species.

Obsoleticeras Bogdanova & I.Mikhailova, gen. nov. - high, quickly rising whorls with oval sections, early
whorls with rare ribs, later with loosening of sculpture. Suture formula with broad shallow elements, loben I,
I

- on the flanks. Type-species: Prodeshayesites obsoletus Casey, 1964, Lower Aptian, fissicostatus Zone,

obsoletus Subzone, England (Surrey). It includes 5 species.

On the whole,

the Deshayesitidae consists of the genera: Turkmeniceras Tovbina, 1962, Deshayesites

Kasansky, 1914, Paradeshayesites Kemper, 1967, Obsoleticeras gen. nov., Dufrenoyia (Burckhardt Ms )
Kilian & Reboul, 1915, Burckhardtites Humphrey, 1949 and probably, Neodeshayesites Casey, 1964 and
Kuntziella Collignon, 1962. The type of suture-development of the last two genera is still unknown.

The Deshayesitidae derived from the Heteroceratidae. The Deshayesitidae has unsteady quinque-lobate
primasuture - VUU1 ID. To the end of the first whorl lobe U1 disappeared and the suture formula is VUID. It
is the same as in the adult Heteroceratidae. Then the fifth lobe arose on the second whorl, but it is the new
lobe I1

in the saddle I/D. In the evolution of the Deshayesitidae the umbilical part of the shell began to

stretch out with involution of whorls and the umbilical part of suture began to incise: VUU'lD - VUID VULI'D - VUII2 :l'D - VUII2!3 :l'D.

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YOLK SAC MORPHOLOGIES IN CEPHALOPOD EMBRYOS
Boletzky, Sigurd v.
Observatoire Oceanologique de Banyuls, F-66651 Banyuls-sur-Mer (E-mail: )

The basic pattern of embryogenesis is rather uniform throughout the class Cephalopoda. Depending on the
specific ovum size, which may vary from less than 1 mm to about 30 mm, the organ-forming part (called
the "embryo cap", or the "embryo proper") of the epibolic gastrula covers a larger (in small eggs) or smaller
part (in large eggs) of the animal hemisphere of the uncleaved yolk mass. The rest of the gastrula forms an
envelope for the yolk that remains outside the embryo cap and becomes the so-called outer yolk sac.

The overall shape of the yolk mass at early embryonic stages varies among systematic groups of
cephalopods, from nearly globular to elongate/oval, to a certain degree also as a function of egg size. At
advanced stages of organogenesis, the outer yolk sac becomes increasingly distinct due to a constriction of
the brachial and cephalic zone of the embryo proper; an exception are some teuthoid squids (especially the
Ommastrephidae), in which the outer yolk sac remains rudimentary. Starting out from a roughly globular
form, the outer yolk sac may subsequently take on a different shape in the embryos of certain taxa.

The portion of the yolk mass lying inside the embryo proper is called the inner yolk sac and yolk neck (the
latter connects the inner with the outer yolk sac). Whereas the yolk neck is a simple strand, which is more
or less strongly compressed by the organs of the head, the inner yolk sac takes on a shape that is only partly
due to simple "moulding" by the surrounding organs. There seems to be a shift of partial pressure between
the outer and the inner yolk sac at late embryonic stages, and the final shape of the inner yolk sac is also
influenced by the morphogenetic processes shaping the whole visceral complex of the embryo. More or less
distinctive morphologies of the inner yolk sac can be recognized; for example, a peculiar "four finger"
pattern appears in sepiolid embryos.

In a strictly functional perspective, the various yolk sac morphologies can be viewed as different modes of
yolk storage during the developmental phase that leads to hatching. In most cephalopods, the newly hatched
young can survive some time without food (from a few days to several weeks, depending on the juvenile
physiology and ecology). Under normal conditions, the embryonic nutriment remaining in the inner yolk sac

of the hatchling is resorbed independently from the onset of digestive processes that are induced by capture
and ingestion of prey. This coexistence of lecithotrophy and active feeding is due to the morphological and
physiological separation of the embryonic and post-embryonic alimentary organs; the actual duration of this
concomitancy is partly conditioned by the volume of the yolk reserve, which in turn depends largely on the
yolk storage capacity of the visceral mass.

This paper views evolutionary variation in developmental morphology of cephalopod "yolk organs" against
the background of some pervasive functional constraints in yolk absorption.

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DWARF

CEPHALOPODS:

CONDITIONS

OF

REPRODUCTION

AT

SMALL

SIZE

Boletzky, Sigurd v.
Observatoire Oceanologique de Banyuls, F-6665I Banyuls-sur-Mer (E-mail: )

The oldest fossils supposed to represent a ccphalopod (Plectronoceras of the Upper Cambrian) are cliambered
shells measuring 15-20 mm in length. Thus the body size was in the same range as the adult size of the
smallest living cephalopods, Idiosepius pygmacus . Only slightly larger are some dwarf sepiids (e. g. Sepia
pulchra , Metasepia spp.), the smallest sepiolids (Sepiola spp), some loliginid squids (Pickfordiateutliis
spp.) and members of several pelagic squid families, some octopods (e. g. Octopus micropyrsus among the
bottom-living forms; dwarf males in the pelagic Argonauta , Tremoctopus , Ocythoe ).

A common biological feature of very small ccphalopod species is a life-span of less than one year, with a
minimum of about 3 months in Idiosepius . (The life span of dwarf males in pelagic octopods is unknown).
The small adult size of the females, along with their short life-span, limits individual fecundity. The
adaptive responses to this limitation vary widely among species or groups; the most striking difference
appears in the respective egg sizes. In Idiosepius pygmacus, the ovum measures about 1 mm in diameter,
whereas in Octopus micropyrsus , it mesures 8-10 mm in length, 4-5 nun in width. In both cases, however,
maturation and release of eggs is prolonged, i. e. this terminal reproduction tends to multiple spawning .

During embryonic development, a very large part of the yolk mass stored in the ovum is used for the
production of embryonic cell complexes and for the subsequent formation of functional organs. But some
yolk remains in the so-called iimer yolk sac; its volume is a function of (1) the speed of yolk absorption
during late embryonic stages, and (2) the point of time when hatching actually occurs. The only species in
which the young hatch without a yolk reserve are medium-sized octopods producing very large eggs
(Octopus maya , Eledonc moschata ) . Hatclilings of E. moschata can survive for 2 to 3 weeks without food,
i. e. the early juvenile metabolism can function according to the adult "emergency program" (using muscular
proteins as an energy source). This is an exceptional capacity for an early juvenile metabolism.

Adult size may be attained rapidly when hatclilings are relatively large due to strong embryonic growtli based
on a large amount of yolk in the egg (e. g. Octopus micropyrsus ). However, Idiosepius shows that even
small eggs giving rise to small hatclilings permit rapid growtli to adult size, within a very short life-span.

A comparative analysis of all ccphalopod embryos large and small suggests one generalization:
embryogenetic mode and juvenile design require a minimal ovum size above 0.5 mm. This generalization is
essential for any hypothesis on the likely reproductive mode of the earliest cephalopods, which were dwarfs.

The focus of this paper is on biological aspects of size limitations in the evolution of cephalopods, with
special emphasis on the secondary size reductions that must have occurred, and on their functional limits.

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PARALLEL BIOZONATION IN THE UPPER CALLOVIAN AND THE LOWER OXFORDIAN
BASED ON THE PELTOCERATINAE SUB-FAMILY (AMMONITINA, ASPIDOCERATIDAE)
Bonnot, Alain, Marchand. Didier. Courville, Philippe
Centre des Sciences de la Terre de l'Universite de Bourgogne, UMR 5561, 6, bd Gabriel
F-21000Dijon
e-mail:

Biozonation in the Upper Callovian and in the Lower Oxfordian
Ammonite biozonation in the Upper Callovian is based on two standard scales: (1) in the Sub-Boreal
province two sub-families of Boreal origin are utilised (Kosmoceratinae and Cardioceratinae); (2) in the SubMediterranean province the scale is mixed, with some index fossils of Tethyan origin (Reineckeiinae,
Pseudoperisphinctinae or Peltoceratinae) while others are of Boreal origin (Cardioceratinae). In the Lower
Oxfordian, zonation for the Sub-Boreal province is reliant, barring exceptions, on the Cardioceratinae. As
these may be rare or absent in the Mediterranean province, it is essential to have a scale based exclusively on
taxa of Tethyan origin. The first subdivisions based either on the Perisphinctidae or on the
Taramelliceratinae are incomplete.
Interest of the Peltoceratinae subfamily
Revision of the Aspidoceratidae family (Bonnot 1995, Courville & Bonnot 1998) shows that the

Peltoceratinae allow a more complete subdivision. This sub-family occurs in the Sub-Mediterranean province
right at the base of the Upper Callovian and remains comparatively abundant through to the top of the Lower
Oxfordian. The Peltoceratinae are well defined by their ontogenesis and sexual dimorphism. Three characters
constitute reliable criteria for identification: (1) the medium position of the rib bifurcation point on the sides
(macroconchs and microconchs), (2) presence or absence of an intermediate ornemental stage between
costulate and tuberculate stages (macroconchs), and (3) the morphology of the latero-ventral tubercles
(macroconchs). These characters have made it possible to define or redefine twelve palaeospecies, with
macroconch and microconch, divided into two genera (Pejtpcgras and Peltoceratoides").
Conclusion
Their relative abundance, their wide geographical distribution and their rapid evolution mean that some
species of Peltoceratinae can be used as index fossils for a biochronological scale at the resolution of the subzone or even the horizon. We propose a scale based on Peltoceratinae alone in the Upper Callovian and the
Lower Oxfordian. This scale is particularly useful when index fossils of Boreal origin are rare or absent.

References
BONNOT A. - 1995 - Les Aspidoceratidae d'Europe occidentale au Callovien supfrieur et ä l'Oxfordien
infeYieur. These Universite Dijon, Inedit: 487 p.
COURVILLE P. & BONNOT A. - 1998 - Faunes ammonitiques et biochronologie de la zone k Athleta et
de la base de la zone ä Lamberti (Callovien superieur) de la Cote de Meuse (France). Interets des faunes
nouvelles d'Aspidoceratidae. Revue de Pal., 17(2) : 307-346.
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